Soil Environmental Factors Drive Seed Density Across Vegetation Types on the Tibetan Plateau
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Plant Soil DOI 10.1007/s11104-017-3348-0 REGULAR ARTICLE Soil environmental factors drive seed density across vegetation types on the Tibetan Plateau Miaojun Ma & James W. Dalling & Zhen Ma & Xianhui Zhou Received: 7 March 2017 /Accepted: 13 July 2017 # Springer International Publishing AG 2017 Abstract Methods Here we collected 1026 soil seed bank sam- Background and aims Soil seed banks are an impor- ples and sampled 171 aboveground vegetation quadrats tant source of seedling recruits in grassland ecosys- in 57 sites representing six distinct grass-dominated tems, particularly following large-scale disturbance. vegetation types present at high elevation on the Tibetan Nonetheless, the relative importance of above- Plateau, China. To understand processes affecting seed ground plant communities versus below-ground pro- banks at these sites we examined the associations of soil cesses in maintaining these seed banks is poorly environmental variables with community composition, understood. density and species richness. Results We found significant differences in species composition between the seed bank and standing Responsible Editor: Jeffrey Walck. vegetation in each of the vegetation types, whereas Electronic supplementary material The online version of this soil seed banks were much more similar to each article (doi:10.1007/s11104-017-3348-0) contains supplementary other. Nonetheless, seed bank composition was sig- material, which is available to authorized users. nificantly associated with soil moisture, pH, and M. Ma (*) : Z. Ma : X. Zhou available nitrogen (all p < 0.05). Seed density was State Key Laboratory of Grassland and Agro-ecosystems, School significantly negatively correlated with soil pH and of Life Sciences, Lanzhou University, Lanzhou, Gansu 730000, positively correlated with soil moisture and soil or- China e-mail: [email protected] ganic matter. Multiple regression analysis showed that for seed bank density a model including pH Z. Ma alone had the lowest AIC value. e-mail: [email protected] Conclusions Soil conditions influence not only seed inputs but also potentially seed survival in the soil, X. Zhou e-mail: [email protected] this study present a framework which supply a plau- sible explanation on effect of soil environment fac- tors (chemical and physical factors) in the formation J. W. Dalling of species composition of soil seed bank. Future Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL 61801, USA work should isolate the direct and indirect effects e-mail: [email protected] of soil chemistry on seed persistence using seed burial experiments. Z. Ma Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, Chinese Academy of Keywords pH . Plant community. Similarity. Soil Sciences, Xining, Qinghai 810008, China environmental factor. Soil moisture . Soil seed bank Plant Soil Introduction Lauber et al. 2008). In particular, the composition of fungal communities can be sensitive to variability in Soil seed banks represent a critical source of recruits soil nutrient availability (Lauber et al. 2008). In for many plant populations that occupy temporally contrast, the composition of soil bacterial communi- variable environments (Venable and Brown 1988). ties is more strongly related to soil pH (Fierer and For some species, seed persistence has been shown Jackson 2006; Hartman et al. 2008; Lauber et al. to be influenced by biotic (e.g. Schafer and Kotanen 2009), potentially reflecting a narrower optimal pH 2003, 2004; Dalling et al. 2011;Mordecai2012)or range for bacterial than fungal growth (Rousk et al. abiotic conditions in the soil (e.g. Pakeman et al. 2010). As yet, however, the effects of soil pH on 2012;Abedietal.2014; Basto et al. 2015a). Deter- grassland seed bank richness and composition has mining how these environmental factors influence received little attention (Basto et al. 2015a), al- seed survival and germination is therefore critical though recent work in acidic and calcareous grass- to understanding the population and community dy- lands in the UK suggests that seed persistence, as namics of seed bank forming species, and has im- well as seed bank richness and density decrease with plications for how seed banks are managed in resto- increasing pH and is potentially mediated by patho- ration ecology (Khurana and Singh 2001). gens (Basto et al. 2015b). Little is known about how In this study, we have proposed a conceptual soil environmental factors potentially affect the spe- model of the direct and indirect effect of soil envi- cies composition, density and richness of soil seed ronmental factors on seed survival and species com- banks across the biomes. position of soil seed bank through have direct effects On the Tibetan plateau diverse grassland and on aboveground plant community and belowground meadow ecosystems have been recognized that re- soil microbes (Fig. 1). Soil seed bank is a balance of flect variation in elevation, moisture regime and soil seed input and seed output (Fenner 1985). Plant properties (Qi et al. 2015). However, no study cur- communities have a direct effect (seed input) on soil rently exists to identify common trends in above- seed bank through seed dispersal. Soil microbes also ground vegetation and seed bank composition across have a direct effect to soil seed bank through path- these different vegetation types. While the effects of ogens invasion. Soil seed bank experiences a com- environmental factors on the seed bank of species plicated ecological process in the soil. However, this present in above-ground vegetation (transient seed process is often neglected. What role does soil en- bank) may be primarily determined via effects on vironment play during soil seed bank formation and local seed production (Fig. 1), environmental effects function is an unresolved scientific issue. on species absent from aboveground vegetation Abiotic environmental conditions in the soil may (persistent seed bank) may instead primarily repre- not only have a direct impact on ungerminated seeds, sent direct effects on seeds mediated over long pe- but also influence seed survival via effects on seed riods by the soil physical and chemical environment, predators and pathogens (Fig. 1). In particular, fungal and by the action of seed predators and pathogens. pathogens have been shown to reduce seed survival The objectives of this paper were therefore: (1) in many habitats (e.g. Dalling et al. 1998; Schafer To determine how the density, richness and compo- and Kotanen 2003, 2004;O’Hanlon-Manners and sitional similarity of seed banks compares to above- Kotanen 2006), with the activity of soil fungi linked ground communities across six Tibetan plateau veg- to soil moisture and other soil characteristics (Blaney etation types, and (2) To determine how seed bank and Kotanen 2001; Schafer and Kotanen 2004; density, richness and species composition varies in Mordecai 2012). As a consequence, wetter habitats relation to soil moisture and soil chemical variables may have lower seed survival at least in part due to (pH, nitrogen and phosphorus availability, and soil the effect of fungal pathogens, while arid habitats organic carbon). Specifically, we hypothesized, that have longer-lived seed banks (Mordecai 2012;Kaiser the richness and density of the persistent seed bank and Pirhofer-Walzl 2015). would be negatively affected by increasing soil Previous work has also shown that soil chemical moisture and pH reflecting increasing losses of variables are often correlated with soil microbial seeds under conditions that favor seed infection communities across space (e.g. Nilsson et al. 2007; and decay by soil microbial communities. Plant Soil Fig. 1 Conceptual model of direct and indirect effects of soil Plant community environmental factors on soil seed banks Seed dispersal Direct effect Indirect effect Direct effect Soil environmental factors Soil seed bank (Soil moisture, pH, SOM, TN, TP, AN, AP…) Direct effect Indirect effect Direct effect Pathogen infection Soil microbes Materials and methods 100 m × 100 m plots at least 1000 m apart were selected at random. Each plot was considered an independent Study sites spatial replicate in each site. We randomly chose three subplots (10 m × 10 m) in each plot, and 10 cylindrical The study was carried out in the northeastern part of soil cores (3.6 cm diameter) were taken randomly from Tibetan Plateau in Gansu Province, P.R China (101°06′- each subplot for a total sample area of 0.092 m2 and the 103°33′E, 33°22′-35°24′N, about 37,000 km2,altitude total volume was 0.0092 m3 per site. The soil cores were 2339–4038 m a.s.l). We chose six vegetation types divided into two fractions: the upper layer (0–5cm (Table S1 in Appendix S1, Table 1), classified as AM depth), and lower layer (5–10 cm depth). Further, the (alpine meadow, 27 sites), ASM (alpine scrub meadow, 10 cores from each depth were then pooled to generate 8 sites), SM (swamp meadow, 7 sites), SAM (subalpine one representative sample per subplot. In total, there meadow, 6 sites), UG (upland grassland, 5 sites), and were 18 samples from each site, 57 sites and 1026 soil FEM (forest edge meadow, 4 sites). The different veg- samples. All the soil samples were transported to the etation types have been distinguished based on species Hezuo Research Station and placed under a north-facing composition, and the identity of dominant species (Wu window of direct exposure at ambient room tempera- 1980). To examine the characteristics of the soil seed tures for 15 days to allow them to dry. Then, samples bank community, a germination experiment was con- were kept dry in a dark storeroom of Hezuo Research ducted in the Research Station of Alpine Meadow and StationwithnoheatuntilMay2010. Wetland Ecosystems of Lanzhou University (Hezuo Branch), Gansu Province of China (N34°55′, Characterization of the germinable soil seed bank E102°53′, altitude 2900 m a.s.l), which was also located on the northeastern part of Tibetan Plateau.